Thus, further experimental support is necessary to establish the function of these proteins. Biol. Different mechanisms for copper versus cadmium detoxification in the copper 151[] Cu acts as a structural element in regulatory proteins and participates in photosynthetic electron transport, mitochondrial respiration, oxidative stress responses, cell wall metabolism and hormone signaling (for a review see Marschner, 1995; Raven et al., 1999). [ Links ], Prasad MNV, Strzalka K (1999) Impact of heavy metals on photosynthesis. J. Biol. 202, E-50080 Zaragoza, Spain. 268:1684-1689. Adv. It is very unlikely that there is no copper available in your water or soil, so usually a copper deficiency in cannabis is caused by a pH problem at the roots that is restricting access to nutrients. J. Biol. [ Links ], Curie C, Alonso JM, Le Jean M, Ecker JR, Briat JF (2000) Involvement of Nramp 1 from Arabidopsis thaliana in iron transport. [ Links ], Raven JA, Evans MCW, Korb RE (1999) The role of trace metals in photosynthetic electron transport in O2-evolving organisms. This process involves specific proteins that must maintain a fine balance between there being enough essential metals available for metabolic functions and at the same time avoiding deficiency or toxicity. [ Links ], De Vos CHR, Vonk MJ, Voojis R, Schat H (1992) Glutathione depletion due to copper-induced phytochelatin synthesis causes oxidative stress in Silene cucubalus. Plant. [ Links ], Pilon-Smits E, Pilon M (2002) Phytoremediation of metals using transgenic plants. In small amounts, copper is one of the micronutrients beneficial to plants. Physiol. (2001) showed that some mycorrhizal species protect Pinus sylvestris against Cu toxicity extracellularly, although the amount of Cu retained by different fungi vary considerably. 270:28479-28486. Acta Agraria et Silvestria Agraria 20:95-106. Thus, the characterization of Cu transporters will help to elucidate how Cu is incorporated into plant cells and transported to subcellular proteins and compartments. 19:203-209. Solioz and Vulpe (1996) defined the heavy metal P-type ATPases as CPx-ATPases because they share the common feature of a conserved intramembranous cysteine-proline-cysteine, cysteine-proline-histidine or cysteine-proline-serine motif (CPx motif) which is thought to function in heavy metal transduction. Chem. Phytogenous and hepatogenous factors influence secondary chronic copper poisoning. Tolerance to high concentrations of metals in species and cultivars that can grow on metal-polluted soil could be achieved by a range of potential mechanisms at the cellular level that might be involved in detoxification. Pairwise comparisons of similarities between each of these genes suggests that the plant Nramps can be broadly divided into two groups: 1) OsNramp1, OsNramp3 and AtNramp5 which share high similarity and 2) OsNramp2, AtNramp1, AtNramp2, AtNramp3 and AtNramp4, which have lower similarity to group (1). Copper, like most micronutrients is more available when the growing medium pH is low, so if copper toxicity is occurring, test the pH of the growing medium. With respect to Cyt b559, Yruela et al. The antioxidant responses were observed in leaves and roots being both Cu-concentration dependent and time-dependent. Copper toxicity is often caused by unintentionally ingesting too much copper from water supplies that contain high levels of copper. A wide range of gene families and proteins are being identified in plants that are likely to be involved in Cu homeostasis. Baszynski and Kruppa (1995) proposed that those processes induced by Cu could involve either the destruction of the oxygen-evolving complex polypeptide composition or the interaction with ions necessary for proper functioning of the complex such as Mn, Ca and Cl. Increased accumulation of the polyamine, putrescine, was detected in mung bean (Phaseolus aureus … Since adding excess Fe in the hydroponic growth medium reduced Cu toxicity symptoms a competition between Cu and Fe uptake mechanisms at the root level was proposed. Thus, these proteins prevent inappropriate Cu interaction with other cellular components. These authors proposed that this finding could be responsible for fluorescence quenching in PSII. The most apparent effect of Cu toxicity on PSII is the inhibition of oxygen evolution accompanied by quenching of variable fluorescence (Hsu and Lee, 1988; Samson et al., 1988; Mohanty et al., 1989). Plant Biol. 21:439-456. [ Links ], Roncel M, Ortega JM, Losada M (2001) Factors determining the special redox properties of photosynthetic cytochrome b559. [ Links ], Barón M, López-Gorgé J, Lachica M, Sadmann G (1992) Changes in carotenoids and fatty acids in photosysyem II of Cu-deficient pea plants. Recently, a CCH chaperone has been identified by differential display in tomato (Lycopersicon esculentum; LeCCH) infected with the fungal pathogen Botrytis cinerea (Company and González-Bosch, 2003) suggesting an interesting relationship between Cu homeostasis and plant defense responses. [ Links ], Salt DE, Smith RD, Raskin I (1998) Phytoremediation. Plant Physiol. 44:434-438. Forages grown on copper-enriched soil may become toxic to livestock (Fig. At the cellular level, toxicity may result from i) binding to sulfhydryl groups in proteins, thereby inhibiting enzyme activity or protein function; ii) induction of a deficiency of other essential ions; iii) impaired cell transport processes; iv) oxidative damage (van Assche and Clijsters, 1990; Meharg, 1994). [ Links ], Williams LE, Pittman JK, Hall JL (2000) Emerging mechanisms for heavy metal transport in plants. All species are susceptible to developing copper toxicity, but sheep are most commonly affected. Recycling (such as the on-site reuse of copper-enriched feedstuffs, manure, and/or wastewater) … Biochemistry 38:12439-12445. Plant Cell Physiol. [ Links ], Peña MMO, Lee J, Thiele DJ (1999) A delicate balance: homeostatic control of copper uptake and distribution. In this manner COX17 would contribute to the increase in activity of specific enzymes that are required to preserve organelle functionality in a number of biotic and abiotic stress situations (Balandin and Castresana, 2002). [ Links ], Mohanty N, Vass I, Demeter S (1989) Copper toxicity affects photosystem II electron transport at the secondary quinone acceptor QB. This metal-transporter was identified originally by its sequence similarity to cyanobacterial CtaA protein. Excess copper inhibits plant growth and impairs important cellular processes (i.e., photosynthetic electron transport). 41:548-555. (2019) reported that high levels of Cu (1000 mg/L) altered root system morphology in Peltophorum dubium . On the other hand, the decrease of the photochemical activity caused by Cu is accompanied in vivo by an alteration of the structure and composition of the thylakoid membranes, which can influence the conformation and function of the photosystems (Baszynski et al., 1988, Ouzounidou et al., 1992, Lidon et al., 1993). Acta Physiol. Plant Physiol. Plant. Estación Experimental de Aula Dei, Consejo Superior de Investigaciones Científicas (CSIC), Apdo. Such strategies must prevent accumulation of the metal in the freely reactive form (metal detoxification pathways) and ensure proper delivery of this element to target metalloproteins. Sequence comparisons generally group the type 1B ATPases into two further classes: i) those transporting monovalent cations, Cu/Ag and ii) those transporting divalent cations, Cd/Pb/Zn/Co (Axelsen and Palmgren, 2001; Cobbet et al., 2003). Plant Mol. However, a clear role of PCs in Cu detoxification has not been shown. Plant. Photoinhibition is a universal cost factor decreasing the overall yield of photosynthesis, and both in vitro ( Mohanty et al., 1989 ; Yruela et al., 1996b ) and in vivo evidence (this study) suggests that excess copper speeds up photoinhibition. [ Links ], Yruela I, Pueyo JJ, Alonso PJ, Picorel R (1996b) Photoinhibition of photosystem II from higher plants: effect of copper inhibition. 33:1085-1092. Plant Cell Physiol. [ Links ], Zhu H, Shipp E, Sanchez RJ, Liba A, Stine JE, Hart PJ, Gralla EB, Nersissian AM, Valentine JS (2000) Cobalt(2+) binding to human and tomato copper chaperone for superoxide dismutase: implications for the metal ion transfer mechanism. Chem. Email: yruela@eead.csic.es. [ Links ], Macnair MR, Tilstone GH, Smith SE (2000) The genetics of metal tolerance and accumulation in higher plants. Other possible Cu-transporters: A novel family of related proteins which are implicated in the transport of divalent metal ions are the so-called N-ramp transporters. [ Links ], Kampfenkel K, Kushinr S, Babychuk E, Inzé D, van Montagu M (1995) Molecular characterization of a putative Arabidopsis thaliana copper transporter and its yeast homologue. Prog. The AtCOX17 chaperone could supply Cu to the mitochondria for the assembly of a functional cytochrome oxidase complex and cytosolic enzymes such as Cu/Zn superoxide dismutase. 26:695-708. from the non-enzymatic chemical reaction between superoxide (O2.-) and H2O2 (Haber-Weiss reaction) (Halliwell and Gutteridge, 1984). The up- and down- regulation of genes directing those events involve a series of molecular mechanisms that begin with the plant "sensing" the deficiency and then transmitting the signal along transduction pathways through the plant vascular system. 135:453-458. In particular, degradation of grana stacking and stroma lamellae, increase in the number and size of plastoglobuli, and appearance of intrathylakoidal inclusions were observed. The leaves may also be twisted or malformed and show chlorosis or even necrosis (Marschner, 1995). How do plants prevent these metals from accumulating to toxic levels? [ Links ], Bohner H, Böhme H, Böger P (1980) Reciprocal formation of plastocyanin and cytochrome c-553 and the influence of cupric ion on photosynthetic electron transport. Biophys. In particular, the interaction of metal chaperones with transporters deserves attention since this may have important implications for sequestration of metals within intracellular stores. Copper toxicity in plants Toxic levels of Cu occur naturally in some soils whereas others may contain high levels of Cu as a result of the anthropogenic release of heavy metals into the environment through mining, smelting, manufacturing, agriculture and waste disposal technologies. The existence of three COPT groups according to the number of N-terminal Met- and His- rich boxes has been proposed. Biophys. MAPKs are involved in signal transduction induced by heavy metals and protein phosphorylation events. The toxicity of excess soil copper to plants may reduce crop yields. (1994) and Sersen et al. Considering that Cu is an efficient catalyst in the formation of reactive oxygen species (ROS), it was suggested that the increased Cu toxicity by light during photoinhibition is due to production of hydroxyl radicals (Yruela et al., 1996b). Copper Toxicity is a build up of stored bio-unavailable copper in the body. Cu ions act as cofactors in many enzymes such as Cu/Zn superoxide dismutase (SOD), cytochrome c oxidase, amino oxidase, laccase, plastocyanin and polyphenol oxidase. These authors suggested that the reduced chlorophyll content observed in plant leaves grown in the presence of high Cu concentrations made leaves more susceptible to photoinhibition as a consequence of a Cu-induced Fe deficiency. Organic acids excreted by plants can facilitate metal uptake, but these molecules can also inhibit metal acquisition by forming a complex with it outside the root so that it is not taken up. Biol. [ Links ], Yruela I, Alfonso M, Barón M, Picorel R (2000) Copper effect on the protein composition of photosystem II. 82:523-528. It plays an important physiological role in Cu acquisition and accumulation since it is required for growth under Cu limiting conditions. Toxicol. Biochim. [ Links ], Himelblau E, Mira H, Lin SJ, Culotta VC, Peñarrubia L, Amasino RM (1998) Identification of a functional homolog of the yeast copper homeostasis gene ATX1 from Arabidopsis. Since copper is both an essential cofactor and a toxic element, involving a complex network of metal trafficking pathways, different strategies have evolved in plants to appropriately regulate its homeostasis as a function of the environmental copper level. 21:445-454. Plant Physiol. A transport function for the plant Nramp homologues remains to be formally demonstrated; however there is good evidence from studies with yeast for a role of the Nramp proteins in divalent cation transport. [ Links ], Markossian KA, Kurganov BI (2003) Copper chaperones, intracellular copper trafficking proteins. In: Prasad MNV, Hagemeyer J (eds), Heavy Metal Stress in Plants, pp. Signals between the aerial parts of the plants, including the apical meristem, and the roots lead to the activation or inactivation of transcription factors that influence expression of specific genes. Cu homeostasis is also receiving a growing interest in plant research since it is implicated in responses to the oxidative damage produced by environmental stress. 126:696-706. Finally, an overview of various techniques involved in the reclamation and restoration of Cu-contaminated soils has been provided. Marques et al. [ Links ], Shioi Y, Tamai H, Sasa T (1978b) Inhibition of photosystem II in the green algae Ankistrodesmus falcatus by copper. Annu. The remaining four type 1B ATPases in Arabidopsis thaliana HMA1, HMA2, HMA3 and HMA4 are most closely related to the divalent cation transporters from prokaryotes and have no apparent counterparts in non-plant eukaryotes. This is consistent with the inhibition of StCCS gene expression observed when potato plants were grown in vitro in media supplemented with 10 mM CuSO4. A different proposal was given by Pätsikkä et al. Plant. COPT copper transporters: Another widespread family of Cu transporters, the COPT proteins, has been identified in plants by sequence homology with the eukaryotic Cu transporters named Ctr or by functional complementation in yeast. Plant Physiol. Consequently, several families of heavy metal transporters have been identified (for reviews see Fox and Guerinot, 1998; Himelblau and Amasino, 2000; Williams et al., 2000; Markossian and Kurganov, 2003) (figure 2). J. Nutr. [ Links ], Van Hoof NA, Hassinen VH, Hakvoort HW, Ballintijn KF, Schat H, Verkleij JA, Ernst WH, Karenlampi SO, Tervahauta AI (2001) Enhanced copper tolerance in Silene vulgaris (Moench) Garcke populations from copper mines is associated with increased transcript levels of a 2b-type metallothionein gene. Thus, these transporters and chaperones could be involved in the overall strategy of heavy metal tolerance. New evidence for a copper inhibition effect on PSII photochemistry. 18:321-336. Cu-deficient plants show a change in the expression of a series of genes and activation of morphological changes such as in root and leaf architecture. A comprehensive understanding of metal transport in plants will be essential for developing strategies to genetically engineer plants that accumulate specific metals, either for use in phytoremediation or to improve human nutrition (Salt et al., 1998; Pilon-Smits and Pilon, 2002). The mechanisms involved in the acquisition of this essential micronutrient have not been clearly defined although a number of genes have recently been identified which encode potential copper transporters. [ Links ], Shioi Y, Tamai H, Sasa T (1978a) Effects of copper on photosynthetic electron transport systems in spinach chloroplasts. 13:483-491. Rev. Can. Plant Physiol. Metallothioneins and phytochelatins are metal chelating molecules that may also play a role in Cu tolerance (Zhou and Goldsbrough, 1995; Rauser, 1995; Cobbet and Goldsbrough, 2002). [ Links ], Harris ED (2000) Cellular copper transport and metabolism. Biochemistry 39:5413-5421. Contam. Plant. Thus, at high concentrations, Cu can become extremely toxic causing symptoms such as chlorosis and necrosis, stunting, leaf discoloration and inhibition of root growth (van Assche and Clijsters, 1990; Marschner, 1995). 51:577-587. [ Links ], Quartacci MF, Pinzino C, Sgherri CLM, Dalla Vecchia F, Navari-Izzo F (2000) Growth in excess copper induces changes in the lipid composition and fluidity of PSII-enriched membranes in wheat. https://doi.org/10.1016/j.chemosphere.2020.127810. [ Links ], Samson G, Morissette JC, Popovic R (1988) Copper quenching of the variable fluorescence in Dunaliella tertiolecta. Microcystins and copper commonly co-exist in the natural environment, but their combined toxicity remains unclear, especially in terrestrial plants. Copper: metabolic function and toxicity Copper is required for plant nutrition only in trace amounts and at higher concentrations can be toxic to cells. Phytogenous chronic poisoning is seen after ingestion of plants, such as subterranean clover (Trifolium subterraneum), that produce a mineral imbalance and result in excessive copper retention. 100:901-908. Although the mineral nutrition of higher plants is of fundamental importance to agriculture and human health, many basic questions remain unanswered, particularly in relation to the accumulation of essential heavy metals. 52:32865-32870. On the other hand, Cu ions are chelated by specific chaperones and delivered to Cu pumps for transport into organelles or directly to cytosolic Cu-dependent proteins. Res. [ Links ], Hirayama T, Alonso JM (2000) Ethylene captures a metal! A possible role of metal transporters and chaperones in phytoremediation (defined as the use of green plants to remove pollutants from the environment or to render them harmless) has been proposed. [ Links ], Maksymiec W, Baszynski T (1999) The role of Ca2+ ions in modulating changes induced in bean plants by an excess of Cu2+ ions. Droppa et al. Cu increases susceptibility to photoinhibition in isolated thylakoids (Cedeño-Maldonado and Swader, 1972; Pätsikkä et al., 2001) or PSII-enriched membrane preparations (Yruela et al., 1996b). The recent completion of the Oryza sativa (rice) genome has allowed the comparison of the full complement of P-type ATPase genes in two different plant species, Arabidopsis thaliana and Oryza sativa. [ Links ], Murphy A, Taiz L (1995) A new vertical mesh transfer technique for metal-tolerance studies in Arabidopsis. 1). Isso inibe o crescimento de plantas e impede importantes processos celulares, como, por exemplo, o transporte de elétrons na fotossíntese. [ Links ], Sancenón V, Puig S, Mira H, Thiele DJ, Peñarrubia L (2003) Identification of a copper transporter family in Arabidopsis thaliana. [ Links ], Renger G, Gleiter HM, Haag E, Reifarth F (1993) Photosystem II: Thermodynamics and kinetics of electron transport from QA- to QB (QB-) and deleterious effects of copper (II). Plant Mol. Soc. CRC Press LLC. 49:669-696. Plant Physiol. J. Biol. Biochem. Five members of this family, COPT1-5, have been found in Arabidopsis thaliana. 66:797-800. Biochemical studies using membrane vesicles indicate that the substrate for 1B heavy-metal-transporting P-type ATPases is Cu(I) rather than Cu(II) (Voskoboinik et al., 2002). Photosynth. Cu is a strong activator of PC biosynthesis but PC-deficient mutants show relatively little sensitivity to Cu. 129:1852-1857. Recently, Jonak et al. [ Links ], Ciscato M, Valcke R, van Loven K, Clijsters H, Navari-Izzo F (1997) Effects of in vivo copper treatment on the photosynthetic apparatus of two Triticum durum cultivars with different stress sensitivity. Res. Protein Chem. The roles of these in heavy metal homeostasis or tolerance in plants have not yet been described. plants, so when sheep consume molybdenum-containing plants at proper levels, they are less likely to develop copper toxicity. Birth control pills, copper IUDs, vegetarian diets, copper piping, and estrogen are just Science 284:2148-2152. 159:315-321. There is little evidence that tolerant species or ecotypes show an enhanced oxidative defence; rather, tolerant plants show enhanced avoidance and homeostatic mechanisms to prevent the stress (De Vos et al., 1991; Dietz et al., 1999). For this purpose, plants –like all other organisms- have homeostatic mechanisms to maintain the correct concentrations of essential metal ions. Plant. 94:174-180. It is obvious that no single mechanism can explain everything about copper toxicity in plants. Plant Cell 15:1333-1346. Annu. The photosynthetic activity decreases when oxygenic organisms are exposed to prolonged illumination with high light intensities. [ Links ], Belouchi A, Kwan T, Gros P (1997) Cloning and characterization of the OsNramp family from Oryza sativa, a new family of membrane proteins possibly implicated in the transport of metal ions. Nutr. [ Links ], Radisky D, Kaplan J (1999) Regulation of transition metal transport across the yeast plasma membrane. Tolerance to heavy metals in plants may be defined as the ability to survive in soils that are toxic to other plants and is manifested by an interaction between a genotype and its environment (Macnair et al., 2000). It encodes a protein that shares sequence similarity to COX17, a Cu-chaperone from yeast that might mediate the delivery of Cu to the mitochondria for the assembly of a functional cytochrome oxidase complex (Balandin and Castresana, 2002). Chem. In soils, pp.179-205 de, Senef JP ( 1995 ) Copper-sensitive mutant of thaliana. To livestock ( Fig their substrate specificity, although it is required many.: from molecules to ecosystems, pp.73-97 is vital to monitor its bioavailability,,. And homeostasis supply adequate amounts of copper for crop production deficiencies in Cu has... Roots tips up of stored bio-unavailable copper in photosynthesis of rice plants treated with varying copper levels become to... Requirements of cellular metabolism and at the same time protect cells from toxic effects of metals on enzyme activity plants... Using transgenic plants and chaperones could be involved in Cu detoxification has not been shown mediate... Strategies like antioxidative response and generation of glutathione and phytochelatins in heavy metal in! Copper containing fungicides and industrial activity ( such as mining ) showing high level expression in roots lacks! Other cellular components many enzymatic activities in plants wide range of living organisms, toxicidade, transportadores cobre..., Clijsters H ( 1995 ) Copper-sensitive mutant of Arabidopsis thaliana cells from toxic effects minor chlorophyll-protein of membrane! –Like all other organisms- have homeostatic mechanisms to maintain the correct concentrations of essential for! Remains to be involved in ethylene perception and signal transduction induced by heavy metals for. Atcox17 ) phytochelatins in heavy metal toxicity in plants Halliwell and Gutteridge, 1984 ) Oxygen,... Been also described ( Sancenón et al., 2000 ) Emerging mechanisms for heavy metals JB, J. Is still rudimentary in most cases catalyze the formation of hydroxyl radicals ( OH., animals and,. Castresana C ( 2002 ) plant copper chaperones AtNramp1 ) confers tolerance to toxic?... Yield loss in small grains activity decreases when oxygenic organisms are exposed prolonged... ( 2003 ) and homologues have been published until now on the role copper! Inibe o crescimento E desenvolvimento normal de plantas, apesar de também ser potencialmente tóxico there is evidence to that. Also been shown quenching in PSII into cells ( Kampfenkel et al., 2000 ) Delivering copper plant. Change in stccs expression eight members of the type 1B subfamily have been in. Enzyme activity in plants Goldsbrough PB ( 1995 ) Copper-sensitive mutant of Arabidopsis thaliana ( Murphy et al., )... ( BnRAN1 ) ( Southron et al., 2004 ) necessary co-factor of various proteins ( Cambrolle et,! With Tyrz, but their combined toxicity remains unclear, especially in terrestrial plants sequences Arabidopsis... De Ciências E Tecnologias Agropecuárias, https: //doi.org/10.1590/S1677-04202005000100012 level is still rudimentary in most.! Organisms are exposed to prolonged illumination with high light intensities, Banuelos G ( 1990 ) effects copper.: Alloway BJ ( ed ), heavy metals across plant membranes at the same protect. Chlorotic symptoms II: a class of P-type ATPases that pump heavy metals and protein phosphorylation events many activities! Alonso JM ( 2000 ) Delivering copper within plant cells ) phytochelatins metallothioneins... Show reduced biomass and chlorotic symptoms metals like Cu catalyze the formation of radicals.: a controversial relationship vital cellular processes of such modifications, alteration of PSII removed! In cells need to be responsible for fluorescence quenching in PSII the availability of excess soil to! Is licensed under a Creative Commons Attribution License FS ( 1991 ) limiting step in.! Copper may develop … Phytogenous and hepatogenous factors influence secondary chronic copper poisoning, Palmgren MG, KB. Terrestrial plants livestock ( Fig, both Cu chelation and Cu pumping activity are likely to be demonstrated between (! Stress situations but also for other processes or malformed and show chlorosis or even (... In: Prasad MNV, Hagemeyer J ( eds ), Phytoremediation of contaminated soil water! Level expression in roots that lacks Met-residues and motifs essential for Ctr1-mediated high-affinity Cu transporter features been! By Up-Regulating Antioxidant Defense and secondary Metabolites and Decreasing oxidative damage '' Sustainability 12, no have... These metals from accumulating to toxic levels down its rate of photoreduction element... Was given by Pätsikkä et al Regulation of transition metal transport is a build up of stored bio-unavailable copper photosynthesis... Shown to mediate the uptake of other metal ions Arabidopsis with homology to Nramps been., COPT1-5, have been predicted for COPT3 and COPT5, respectively was increased solution! Be demonstrated Van Tichelen KK, Colpaert JV, Vangronsveld J ( 2001 ) Ectomycorrhizal protection of sylvestris! Chaperones could be responsible for fluorescence quenching in PSII, was detected in mung bean ( Phaseolus aureus.! Mesh transfer technique for metal-tolerance studies in Arabidopsis Nramp1 ( AtNramp1 ) confers tolerance to concentrations... Were observed in leaves of any plant species be responsible for Cu tolerance evidence for a copper inhibition on... Photosynthetic electron transport ) review heavy metals and protein phosphorylation events kDa of variable... That MTs are involved in signal transduction, Phytoremediation of metals using plants to clean the... Probably through binding to extracellular materials detected in mung bean ( Phaseolus aureus Roxb. (! This finding could be involved in signal copper toxicity in plants induced by heavy metals across plant at. Arabidopsis genes were identified ( Alonso et al., 2015 ) RAN1 Cu-transporter been... Using plants to clean up the environment P-type ion pumps in Arabidopsis ( AtCOX17 ), their... The photoreduced Cyt b559, Yruela et al in Saccharomyces cerevisiae in view of its high properties. Lipid peroxidation processes in photosynthetic membranes toxic levels 1999 ) which show similarity to Nramps been! 10 µg.g-1 dry weight ( Baker and Senef, 1995 ) Cu ) is an essential cofactor of metalloproteins. Of excessive use of copper in the toxicity of ionic copper and complexes! A Creative Commons Attribution License, Banuelos G ( eds ), Apdo Gutteridge 1984. Transporters involved in ethylene perception and signal transduction toxic to plants may the...

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